Phylogeographic analyses of a circumarctic coastal and a boreal lacustrine mysid crustacean, and evidence of fast postglacial mtDNA rates

Phylogeographic structures of two weakly dispersing Mysis sibling species, one with a circumarctic coastal, the other with a boreal lacustrine-Baltic distribution, were studied from mitochondrial cytochrome c oxidase subunit I (COI) gene sequences. Mysis segerstralei showed high overall diversity and little phylogeographic structure across the Arctic, indicating late-glacial dispersal among coastal and lake populations from Alaska, Siberia and the north of Europe. A strongly divergent refugial lineage was however identified in Beringia. The boreal 'glacial relict'Mysis salemaai in turn displayed clear structuring among postglacially isolated Scandinavian lake populations. The inferred pattern of intralake mitochondrial DNA (mtDNA) monophyly in Scandinavia suggested relatively small population sizes and a remarkably fast postglacial mtDNA divergence rate (0.27% per 10 000 years). Nevertheless, the broader phylogeographic pattern did not support distinct eastern and western glacial refugia in Northern Europe, unlike in some other aquatic taxa. In all, the two species comprised three equidistant mitochondrial lineages (approximately 2% divergence), corresponding to M. salemaai, to the bulk of M. segerstralei, and to the Beringian M. segerstralei lineage. The lack of reciprocal monophyly of the two species in respect to their mitochondrial genealogy could indicate postspeciation mitochondrial introgression, also exemplified by an evidently more recent capture of M. segerstralei mitochondria in a Karelian population of M. salemaai. Overall, the data suggest that the continental boreal M. salemaai has a relatively recent ancestry in arctic coastal waters, whereas two other boreal 'glacial relict'Mysis sibling species in Europe (Mysis relicta) and North America (Mysis diluviana) have colonized inland waters much earlier (approximately 8% COI divergence).     

Phylogeography and origin of freshwater populations of tubenose gobies of genus Proterorhinus (Gobiidae: Pisces) in Ponto-Caspian Basin

Representatives of the genus Proterorhinus (tubenose gobies) from the water bodies of the Ponto-Caspian Basin were examined for sequence polymorphism of the mitochondrial DNA fragment containing the cytochrome b (Cyt-b) gene. A total of ten haplotypes were discovered, which formed two groups. Thus, the data obtained indicated the existence of only two taxonomically valid phylogenetic lineages, represented by (1) marine and brackish-water populations of the Black Sea and (2) freshwater populations of the whole Ponto-Caspian Basin, along with the brackish-water population of the Caspian Sea. Based on an analysis of the tubenose goby haplotype distribution patterns, the colonization patterns of this group of fishes (phylogeography) in the freshwater drainages of Ponto-Caspian Basin are examined. It was established that the basin of the ancient Caspian Sea was the major donor area for the formation of the freshwater and brackish-water populations. In the Ponto-Caspian Basin, two centers of the tubenose gobies speciation and distribution are distinguished. One of these centers is associated with the modern northwestern part of the Black Sea, and another one is confined to the Caspian Sea. It is suggested that the modern colonization of the Volga River by tubenose goby occurred from the Caspian Sea.     

Spectral sensitivity differences in two Mysis sibling species (Crustacea, Mysida): Adaptation or phylogenetic constraints?

The variation in eye spectral sensitivities of the closely related mysid species Mysis relicta Lovén, 1862 and Mysis salemaai Audzijonyt? and Väinölä, 2005 was studied in sympatric and allopatric populations from the brackish Baltic Sea and from two lakes representing different light environments. In the Baltic Sea the maximum spectral sensitivity of M. relicta, measured by the electroretinogram (ERG) technique, was shifted by ca 20 nm to longer wavelengths than in M. salemaai (564 and 545 nm, respectively). The spectral sensitivity of M. salemaai was closer to that of marine mysid species, which is consistent with its broader euryhalinity and the presumed longer brackish-water history. The species-specific sensitivities in the Baltic Sea were not affected by regional differences in light environments. In two lake populations of M. relicta, the spectral sensitivity was further shifted by ca 28 nm towards the longer wavelengths compared with the conspecific Baltic Sea populations. The spectral sensitivities in the four M. relicta populations were not correlated to the current light conditions, but rather to the phylogeographic histories and fresh- vs. brackish-water environments. A framework to further explore factors affecting spectral sensitivities in Mysis is suggested.     

Comparative phylogeography of Ponto-Caspian mysid crustaceans: isolation and exchange among dynamic inland sea basins

The distributions of many endemic Ponto-Caspian brackish-water taxa are subdivided among the Black, Azov and Caspian Sea basins and further among river estuaries. Of the two alternative views to explain the distributions, the relict school has claimed Tertiary fragmentation of the once contiguous range by emerging geographical and salinity barriers, whereas the immigration view has suggested recolonization of the westerly populations from the Caspian Sea after extirpation during Late Pleistocene environmental perturbations. A study of mitochondrial (COI) phylogeography of seven mysid crustacean taxa from the genera Limnomysis and Paramysis showed that both scenarios can be valid for different species. Four taxa had distinct lineages related to the major basin subdivision, but the lineage distributions and depths of divergence were not concordant. The data do not support a hypothesis of Late Miocene (10-5 Myr) vicariance; rather, range subdivisions and dispersal from and to the Caspian Sea seem to have occurred at different times throughout the Pleistocene. For example, in Paramysis lacustris each basin had an endemic clade 2-5% diverged from the others, whereas Paramysis kessleri from the southern Caspian and the western Black Sea were nearly identical. Species-specific ecological characteristics such as vagility and salinity tolerance seem to have played important roles in shaping the phylogeographic patterns. The mitochondrial data also suggested recent, human-mediated cryptic invasions of P. lacustris and Limnomysis benedeni from the Caspian to the Sea of Azov basin via the Volga-Don canal. Cryptic species-level subdivisions were recorded in populations attributed to Paramysis baeri, and possibly in P. lacustris.     

Migration and isolation during the turbulent Ponto‐Caspian Pleistocene create high diversity in the crustacean Paramysis lacustris

The Ponto‐Caspian brackish‐water fauna inhabits estuaries and rivers of the Black, Azov and Caspian seas and is fragmented by higher salinity waters and a major interbasin watershed. The fauna is known for the high levels of endemism, complex zoogeographic histories, and as a recent source of successful invasive species. It remains debated whether the Black and Azov Sea brackish‐water populations survived unfavourable Pleistocene conditions in multiple separate refugia or whether the two seas were (repeatedly) recolonized from the Caspian. Using microsatellite and mtDNA markers, we demonstrate deep among‐ and within‐basin subdivisions in a widespread Ponto‐Caspian mysid crustacean Paramysis lacustris. Five genetic clusters were identified, but their relationships did not reflect the geography of the region. The Azov cluster was the most distinct (4–5% COI divergence), despite its geographic position in the corridor between Black and Caspian seas, and may represent a new species. In the northern Black Sea area, the Dnieper cluster was closer to the Caspian cluster than to the neighbouring Danube–Dniester–Bug populations, suggesting separate colonizations of the Black Sea. Overall, the data implied a predominant gene flow from the east to the Black Sea and highlight the importance of Caspian Sea transgressions in facilitating dispersal. Yet, the presence of distinct lineages in the Black Sea points to the persistence of isolated refugial populations that have gained diagnostic differences under presumably high mutation rates and large population sizes. The unfavourable Pleistocene periods in the Black Sea therefore appear to have promoted diversification of the brackish‐water lineages, rather than extirpated them.     

Phylogeography and taxonomic status of trout and salmon from the Ponto‐Caspian drainages,with inferences on European Brown Trout evolution and taxonomy

Current taxonomy of western Eurasian trout leaves a number of questions open; it is not clear to what extent some species are distinct genetically and morphologically. The purpose of this paper was to explore phylogeography and species boundaries in freshwater and anadromous trout from the drainages of the Black and the Caspian Seas (Ponto‐Caspian). We studied morphology and mitochondrial phylogeny, combining samples from the western Caucasus within the potential range of five nominal species of trout that are thought to inhabit this region, and using the sequences available from GenBank. Our results suggest that the genetic diversity of trout in the Ponto‐Caspian region is best explained with the fragmentation of catchments. (1) All trout species from Ponto‐Caspian belong to the same mitochondrial clade, separated from the other trout since the Pleistocene; (2) the southeastern Black Sea area is the most likely place of diversification of this clade, which is closely related to the clades from Anatolia; (3) The species from the Black Sea and the Caspian Sea drainages are monophyletic; (4) except for the basal lineage of the Ponto‐Caspian clade, Salmo rizeensis, all the lineages produce anadromous forms; (5) genetic diversification within the Ponto‐Caspian clade is related to Pleistocene glacial waves; (6) the described morphological differences between the species are not fully diagnostic, and some earlier described differences depend on body size; the differences between freshwater and marine forms exceed those between the different lineages. We suggest a conservative taxonomic approach, using the names S. rizeensis and Salmo labrax for trout from the Black Sea basin and Salmo caspius and Salmo ciscaucasicus for the fish from the Caspian basin.     

Visual pigment absorbance and spectral sensitivity of the <Emphasis Type="Italic">Mysis relicta</Emphasis> species group (Crustacea,Mysida) in different light environments

Visual-pigment absorbance spectra and eye spectral sensitivities were examined in eight populations of opossum shrimp from different light environments. Four Finnish populations, two from the Baltic Sea and two from freshwater lakes, represent Mysis relicta, sensu stricto. The sibling species M. salemaai and M. diluviana are represented by, respectively, two Baltic Sea populations and two populations from freshwater lakes in Idaho, USA. In M. relicta, the visual pigments of the two lake populations were similar (λ_max=554.3±0.8 nm and 556.4±0.4 nm), but significantly red-shifted compared with the sea populations (at 529 and 535 nm) and with M. salemaai (at 521 and 525 nm). All these pigments had only A2 chromophore and the lake/sea difference indicates adaptive evolution of the opsin. In M. diluviana, λ_max varied in the range 505–529 nm and the shapes of spectra suggested varying A1/A2 chromophore proportions, with pure A1 in the 505 nm animals. Eye sensitivity spectra were flatter and peaked at longer wavelengths than the relevant visual-pigment templates, but declined with the same slope beyond ca. 700 nm. The deviations from visual-pigment spectra can be explained by ocular light filters based on three types of identified screening pigments.     

Phylogeography and population history of the least weasel (Mustela nivalis) in the Palearctic based on multilocus analysis

The least weasel (Mustela nivalis) is one of the most widely distributed carnivorans. While previous studies have identified distinct western and eastern mitochondrial DNA (mtDNA) lineages of the species in the western Palearctic, their broader distributions across the Palearctic have remained unknown. To address the broad-scale phylogeographical structure, we expanded the sampling to populations in Eastern Europe, the Urals, the Russian Far East, and Japan, and analyzed the mtDNA control region and cytochrome b, the final intron of the zinc finger protein on Y chromosome (ZFY), and the autosomal agouti signaling protein gene (ASIP). The mtDNA data analysis exposed the previous western lineage (Clade I) but poorly supported assemblage extending across Palearctic, whereas the previous eastern lineage (Clade II) was reconfirmed and limited in the south western part of the Palearctic. The ZFY phylogeny showed a distinctive split that corresponding to the mtDNA lineage split, although less phylogeographical structure was seen in the ASIP variation. Our data concur with the previous inference of the Black Sea–Caspian Sea area having an ancestral character. The Urals region harbored high mitochondrial diversity, with an estimated coalescent time of around 100,000 years, suggesting this could have been a cryptic refugium. Based on the coalescent-based demographic reconstructions, the expansion of Clade I across the Palearctic was remarkably rapid, while Clade II was relatively stable for a longer time. It seems that Clade II has maintained a constant population size in the temperate region, and the expansive Clade I represents adaptation to the cold regions.     

History of aquatic invertebrate invasions in the Caspian Sea

Incorporation of the fossil record and molecular markers into studies of biological invasions provides new historical perspectives on the incidence of natural and human-mediated invasions of nonindigenous species (NIS). Palaeontological, phylogeographic, and molecular evidence suggests that the natural, multiple colonizations of the Caspian basin via transient connections with the Black Sea and other basins played an important role in shaping the diversity of Caspian fauna. Geographically isolated, conspecific Ponto-Caspian lineages that currently inhabit fragmented habitats in the Ponto-Caspian region show limited genetic divergence, implying geologically recent episodes of gene flow between populations during the Pliocene to Pleistocene. Several molluscan lineages in the Caspian Sea may have descended from Lake Pannon stock before the Late Miocene isolation of the Caspian depression, about 5.8 million years ago. Anthropogenic activities during the 20th century were responsible for a 1800-fold increase in the rate of establishment of new aquatic species in the Caspian Sea compared to the preceding two million years of natural colonization. The observed success of NIS invasions during the 20th century was due primarily to human-mediated transport mechanisms, which were dominated by shipping activities (44%). Human-mediated species transfer has been strongly asymmetrical, toward the Volga Delta and Caspian Sea from or through Black and Azov Seas. Global and regional trade, particularly that mediated by commercial ships, provides dispersal opportunities for nonindigenous invertebrates, indicating that future invasions in the Caspian Sea are anticipated.     

Out of the blue: adaptive visual pigment evolution accompanies Amazon invasion

Incursions of marine water into South America during the Miocene prompted colonization of freshwater habitats by ancestrally marine species and present a unique opportunity to study the molecular evolution of adaptations to varying environments. Freshwater and marine environments are distinct in both spectra and average intensities of available light. Here, we investigate the molecular evolution of rhodopsin, the photosensitive pigment in the eye that activates in response to light, in a clade of South American freshwater anchovies derived from a marine ancestral lineage. Using likelihood-based comparative sequence analyses, we found evidence for positive selection in the rhodopsin of freshwater anchovy lineages at sites known to be important for aspects of rhodopsin function such as spectral tuning. No evidence was found for positive selection in marine lineages, nor in three other genes not involved in vision. Our results suggest that an increased rate of rhodopsin evolution was driven by diversification into freshwater habitats, thereby constituting a rare example of molecular evolution mirroring large-scale palaeogeographic events.     

Molecular identification of two strains of third-stage larvae of Contracaecum rudolphii sensu lato (Nematoda: Anisakidae) from fish in Poland

Contracaecum sp. larvae (L3) from fish were identified using nucleotide sequences of the internal transcribed spacers ITS-1 and ITS-2 of the ribosomal DNA. The nematode larvae originated from fish in a freshwater situation (crucian carp Carassius carassius, from Selment Wielki Lake in Mazury, northeastern Poland) and a brackish-water region (Caspian round goby Neogobius melanostomus from the Baltic Sea, Gdafisk Bay at the Polish coast). Two strains (Contracaecum rudolphii A and B) of Contracaecum rudolphii senso lato, a parasite common at the adult stage in fish-eating birds, were identified. In fish from the freshwater site, only the strain temporarily designated C. rudolphii B was identified; in the brackish-water region, both strains were found, suggesting that fish serve as paratenic host for both genotypes. Contracaecum rudolphii sensu lato has been recorded in several species of fish-eating birds in Poland, particularly in the great cormorant, Phalacrocorax carbo, in which the abundance is highest. The results, although based on a restricted number of larvae, suggest that the life cycles of both genotypes can be completed in the Polish region and that at least one of them, C. rudolphii B, can develop both in fresh and brackish water.     

Comparative Analysis of Japanese Three-Spined Stickleback Clades Reveals the Pacific Ocean Lineage Has Adapted to Freshwater Environments while the Japan Sea Has Not

Divergent selection and adaptive divergence can increase phenotypic diversification amongst populations and lineages. Yet adaptive divergence between different environments, habitats or niches does not occur in all lineages. For example, the colonization of freshwater environments by ancestral marine species has triggered adaptive radiation and phenotypic diversification in some taxa but not in others. Studying closely related lineages differing in their ability to diversify is an excellent means of understanding the factors promoting and constraining adaptive evolution. A well-known example of the evolution of increased phenotypic diversification following freshwater colonization is the three-spined stickleback. Two closely related stickleback lineages, the Pacific Ocean and the Japan Sea occur in Japan. However, Japanese freshwater stickleback populations are derived from the Pacific Ocean lineage only, suggesting the Japan Sea lineage is unable to colonize freshwater. Using stable isotope data and trophic morphology, we first show higher rates of phenotypic and ecological diversification between marine and freshwater populations within the Pacific Ocean lineage, confirming adaptive divergence has occurred between the two lineages and within the Pacific Ocean lineage but not in the Japan Sea lineage. We further identified consistent divergence in diet and foraging behaviour between marine forms from each lineage, confirming Pacific Ocean marine sticklebacks, from which all Japanese freshwater populations are derived, are better adapted to freshwater environments than Japan Sea sticklebacks. We suggest adaptive divergence between ancestral marine populations may have played a role in constraining phenotypic diversification and adaptive evolution in Japanese sticklebacks.     

Eye function of Mysidacea (Crustacea) in the northern Baltic Sea

Eye spectral sensitivity, [S(lambda)], was measured in seven northern Baltic mysid species using an electroretinogram technique. Their S(lambda) curves were compared with the spectral distribution of underwater light at their normal habitats. In the littoral species Neomysis integer, Praunus flexuosus and Praunus inermis, the S(lambda) maxima, [S(lambda)(max)], were in the wavelength-bands of 525-535, 505-515 and 520-530 nm respectively. The neoimmigrant species Hemimysis anomala had a S(lambda)(max) around 500 nm and high sensitivity at 393 nm, possibly indicating UV-sensitivity. S(lambda) of the pelagic species Mysis mixta and Mysis relicta sp. II was at about 505-520 nm. M. relicta sp. I from Pojoviken Bay and fresh water humic Lake P??j?rvi had S(lambda)(max) at approximately 550 nm and 570 nm respectively. This is in accordance with a similar long-wavelength shift in light transmittance of the respective waters. The eyes of the latter population were also damaged by strong light. The pontocaspian neoimmigrant H. anomala is clearly adapted to waters transmitting more blue light.     

Historic speciation and recent colonization of Eurasian monkey gobies (Neogobius fluviatilis and N. pallasi) revealed by DNA sequences,microsatellites, and morphology

Aim Hidden diversity within an invasive ‘species’ can mask both invasion pathways and confound management goals. We assessed taxonomic status and population structure of the monkey goby Neogobius fluviatilis across Eurasia, comparing genetic variation across its native and invasive ranges. Location Native populations were analysed within the Black and Caspian Sea basins, including major river drainages (Dnieper, Dniester, Danube, Don and Volga rivers), along with introduced locations within the upper Danube and Vistula river systems. Methods DNA sequences and 10 nuclear microsatellite loci were analysed to test genetic diversity and divergence patterns of native and introduced populations; phylogenetic analysis of mtDNA cytochrome b and nuclear RAG‐1 sequences assessed taxonomic status of Black and Caspian Sea lineages. Multivariate analysis of morphology was used to corroborate phylogenetic patterns. Population genetic structure within each basin was evaluated with mtDNA and microsatellite data using F_ST analogues and Bayesian assignment tests. Results Phylogenetic analysis of mitochondrial and nuclear sequences discerned a pronounced genetic break between monkey gobies in the Black and Caspian Seas, indicating a long‐term species‐level separation dating to c. 3 million years. This pronounced separation further was confirmed from morphological and population genetic divergence. Bayesian inference showed congruent patterns of population structure within the Black Sea basin. Introduced populations in the Danube and Vistula River basins traced to north‐west Black Sea origins, a genetic expansion pattern matching that of other introduced Ponto‐Caspian gobiids. Main conclusions Both genetic and morphological data strongly supported two species of monkey gobies that were formerly identified as subspecies: N. fluviatilis in the Black Sea basin, Don and Volga Rivers, and the Kumo‐Manych Depression, and Neogobius pallasi in the Caspian Sea and Volga River delta. Genetic origins of introduced N. fluviatilis populations indicated a common invasion pathway shared with other introduced Ponto‐Caspian fishes and invertebrates.     

A new Paratethyan genus of the ostracod subfamily Loxoconchinae (Podocopida,Cytheroidea)

A new ostracod genus established in a Loxoconchinae fauna from the Eastern Paratethys, with two species known from the Pliocene–post-Pliocene and Recent of the Caspian Sea and Dnieper–Bug estuary of the Black Sea Basin is described. Comparative morphological analysis of four Loxoconchinae genera from a brackish-water fauna of the Paratethyan basins is performed and the trends in morphological evolution of their hinge structures are revealed.     

ORIGIN AND RECENT ENDEMIC DIVERGENCE OF A CASPIAN MYSIS SPECIES FLOCK WITH AFFINITIES TO THE “GLACIAL RELICT” CRUSTACEANS IN BOREAL LAKES

Aspects of the evolution of intralacustrine species flocks and of the origin of the Arctic or “glacial-relict” zoogeographical element in Eurasian inland waters were elucidated in an allozyme study of the crustacean genus Mysis. This element, of supposedly northern marine ancestry, is represented by vicarious taxa in the deeper parts of the Caspian Sea (an enclosed ancient basin) and in young boreal lakes. The three endemic Caspian Mysis species studied are very close genetically (Nei's D = 0.06), which suggests a recent intrabasin radiation and rapid morphological divergence. This is in contrast to the pattern in postglacial Holarctic boreal lakes, where the Mysis relicta group is represented by a set of morphologically uniform but probably much older sibling species (D = 0.3–0.6). The results provide a parallel to those on the recent diversification of some fish species flocks in ancient freshwater lakes. The situation is, however, unusual in that the Caspian sympatric Mysis flock is pelagic, and conditions promoting speciation through allopatric isolation or spatial segregation by trophic substrate specialization seem implausible. The monophyletic Caspian Mysis clade shows a relatively strong divergence from both the northern lacustrine and the Arctic marine congeners (D = 0.6–1.0); the phylogenetic branching order of these three zoogeographical groups is not conclusively resolved. The results contradict the prevailing hypothesis of a recent Pleistocene origin of the Caspian Arctic element by invasion from Eastern European continental proglacial lakes that drained south to the Caspian basin during the glacial maxima and served as refugia for the boreal lacustrine taxa.     

Molecular characterization of crustacean visual pigments and the evolution of pancrustacean opsins

Investigations of opsin evolution outside of vertebrate systems have long been focused on insect visual pigments, whereas other groups have received little attention. Furthermore, few studies have explicitly investigated the selective influences across all the currently characterized arthropod opsins. In this study, we contribute to the knowledge of crustacean opsins by sequencing 1 opsin gene each from 6 previously uncharacterized crustacean species (Euphausia superba, Homarus gammarus, Archaeomysis grebnitzkii, Holmesimysis costata, Mysis diluviana, and Neomysis americana). Visual pigment spectral absorbances were measured using microspectrophotometry for species not previously characterized (A. grebnitzkii=496 nm, H. costata=512 nm, M. diluviana=501 nm, and N. americana=520 nm). These novel crustacean opsin sequences were included in a phylogenetic analysis with previously characterized arthropod opsin sequences to determine the evolutionary placement relative to the well-established insect spectral clades (long-/middle-/short-wavelength sensitive). Phylogenetic analyses indicate these novel crustacean opsins form a monophyletic clade with previously characterized crayfish opsin sequences and form a sister group to insect middle-/long-wavelength-sensitive opsins. The reconstructed opsin phylogeny and the corresponding spectral data for each sequence were used to investigate selective influences within arthropod, and mainly "pancrustacean," opsin evolution using standard dN/dS ratio methods and more sensitive techniques investigating the amino acid property changes resulting from nonsynonymous replacements in a historical (i.e., phylogenetic) context. Although the conservative dN/dS methods did not detect any selection, 4 amino acid properties (coil tendencies, compressibility, power to be at the middle of an alpha-helix, and refractive index) were found to be influenced by destabilizing positive selection. Ten amino acid sites relating to these properties were found to face the binding pocket, within 4 A of the chromophore and thus have the potential to affect spectral tuning.     

Is salinity an obstacle for biological invasions?

Invasions of freshwater habitats by marine and brackish species have become more frequent in recent years with many of those species originating from the Ponto‐Caspian region. Populations of Ponto‐Caspian species have successfully established in the North and Baltic Seas and their adjoining rivers, as well as in the Great Lakes–St. Lawrence River region. To determine if Ponto‐Caspian taxa more readily acclimatize to and colonize diverse salinity habitats than taxa from other regions, we conducted laboratory experiments on 22 populations of eight gammarid species native to the Ponto‐Caspian, Northern European and Great Lakes–St. Lawrence River regions. In addition, we conducted a literature search to survey salinity ranges of these species worldwide. Finally, to explore evolutionary relationships among examined species and their populations, we sequenced the mitochondrial cytochrome c oxidase subunit I gene (COI) from individuals used for our experiments. Our study revealed that all tested populations tolerate wide ranges of salinity, however, different patterns arose among species from different regions. Ponto‐Caspian taxa showed lower mortality in fresh water, while Northern European taxa showed lower mortality in fully marine conditions. Genetic analyses showed evolutionary divergence among species from different regions. Due to the geological history of the two regions, as well as high tolerance of Ponto‐Caspian species to fresh water, whereas Northern European species are more tolerant of fully marine conditions, we suggest that species originating from the Ponto‐Caspian and Northern European regions may be adapted to freshwater and marine environments, respectively. Consequently, the perception that Ponto‐Caspian species are more successful colonizers might be biased by the fact that areas with highest introduction frequency of NIS (i.e., shipping ports) are environmentally variable habitats which often include freshwater conditions that cannot be tolerated by euryhaline taxa of marine origin.